Regime shifts in the breeding of an Atlantic puffin population

Timing of breeding is a key factor determining the reproductive success in bird populations and known to be affected by climate fluctuations. We investigated the long‐term (1978–2002) relationship between climate and hatching date within a population of Atlantic puffin Fratercula arctica at Røst in the Norwegian Sea. The timing of puffin breeding was found to be influenced by the North Atlantic Oscillation winter index (NAO). We isolated two temporal regimes, one where NAO had a significant effect on hatching date (1978–1986 and 1995–2002) and one where these variables were independent (1987–1994). Hatching date could be modelled using, in addition to NAO, hatching date and food abundance in the preceding breeding season (possibly proxies of parental effort). The models remained significant for regime 1 but not for regime 2. NAO differed between the two regimes suggesting that the shifts were induced by climate change, possibly via its effect on the availability of prey in the preceding year. The novelty of our study is the identification of temporal regimes in the effects of climate within one population.     

Feeding, Ichthyophonus sp. infection, distribution and growth history of Norwegian spring-spawning herring in summer

After its collapse in the late 1960s, 1991 was the first year of systematic herring investigations after the Norwegian Sea stock returned to the traditional summer-feeding areas. Of 11 prey groups three dominated the diet; Calanus finmarchicus, Parathemisto spp. and Sebastes spp. There was a high prevalence of Ichthyophonus sp. at stations in the central and western parts of the Norwegian Sea where the cpue was low. This suggests that infected individuals were unable to return from the feeding to the wintering area of the Norwegian fjords. Highest cpue were recorded between 68–73° N and 5–15° E. Age groups 2, 3 and 8 were most frequently caught and age group 8 (the 1983 year class) dominated the samples. Herring occurred in dense concentrations where C. finmarchicus dominated diet. 0-group fish dominated diet at stations along the coast from 66–73° N and out to c . 300 km off the coast. Mature herring overlapped in distribution with 0-group fish and Sebastes spp. contributed largely to herring diet in this area. The first rich year-class after the collapse, the 1983 year class, had a faster growth than rich year classes from past herring periods, but slower than poor ones.     

A biological sampling problem illustrated by the population structure and growth patterns of Sardinella aurita at Tripoli,Libya

Synopsis Sardinella aurita were sampled from catches of the lampara fishery at Tripoli, Libya during October 1979 through to September 1980. Landings consisted mainly of adult fish with large (mode 21–26 cm total length), relatively fast growing 3–5-group individuals in winter and spring and smaller (mode 14–20 cm total length), slow growing 2- and 3-group fish in summer. Gonad growth commenced in April, when the fish were feeding on zooplankton, and continued at the expense of mesenteric fat until July. It is suggested that the older and bigger fish spawn first, probably after having moved westwards, and that this results in spatial and temporal differences in length at age within the population. There was no consistent difference in growth rate between the sexes, but because first maturity in males tends to occur earlier than in females, there were length distribution differences between the sexes in the youngest (2+) age group as sampled.     

The long-term (1979–2005) effects of the North Atlantic Oscillation on wind-induced wave mixing in Loch Leven (Scotland)

We report on long-term covariation (1979–2005) between indices of the North Atlantic Oscillation (NAO) and wind speed and direction in Loch Leven. The effects of the observed variations in wind speed and direction were combined to produce modelled wave-mixed depths (Zc). Positive correlations were observed between seasonal and annual wind speeds and westerly frequency and indices of the NAO that are in line with general perception: positive NAO was correlated with stronger, more westerly winds and these correlations were strongest in winter and spring. Correlations between NAO and estimates of Zc were strongest in the most westerly exposed site in spring (r ² = 0.701; Zc_spring versus spring NAO index). On average, over a 25-year period Zc was deeper in spring and shallower in summer. Major anomalies from the 25-year seasonal means were observed in 1982, 1979, and 1991. Annual average Zc was low in the late 1970s and early 1980s (shallowest average annual Zc of 1.0 m (1984)), high in the late 1980s and early 1990s (deepest average annual Zc of 1.9 m (1990)) and moderate in recent years (up to 2005). This study has major implications for our understanding of potential climate change drivers and the related responses of shallow lake ecosystems, including alterations to littoral habitat quality and benthic–pelagic coupling.     

Territory occupancy rate of goshawk and gyrfalcon: no evidence of delayed numerical response to grouse numbers

Two recent studies on territory occupancy rates of goshawk Accipiter gentilis and gyrfalcon Falco rusticolus report a 2–3-year-delayed numerical response to grouse numbers, which is a requirement for a hypothesis of predator-generated grouse cycles. The time lags were assumed to reflect the average age of sexual maturity in the raptor species. In southern Norway, however, subadult (two-year-old) goshawk hens occupied only 18–25% of territories where occupancy was not recorded in the preceding year, and there was no significant relationship between the proportion of subadults among recruits and grouse indices two years earlier. We argue that territory occupancy rates are not appropriate indices of total raptor population levels, but rather reflect the proportion of territorial pairs that attempt to nest. Because this depends on the body condition of the hens, fluctuations in other important winter resident prey species (most important for the goshawk) and winter weather (most important for the gyrfalcon) should also be addressed. During 1988–2006, the annual proportion of goshawk territories with recorded nesting attempts in southern Norway was most closely related to the preceding autumn’s population indices of black grouse Tetrao tetrix and mountain hare Lepus timidus, whereas the annual proportion of gyrfalcon territories with observations of falcons or with confirmed breeding attempts in central Norway were best explained by population indices of willow grouse Lagopus lagopus and ptarmigan L. mutus from the previous autumn, and by December temperatures. Hence, our studies do not support the predation hypothesis for grouse cycles.     

Large-scale climatic phenomena and timing of breeding in a local population of the Osprey Pandion haliaetus in southern Finland

The effects of large-scale climatic phenomena on the timing of breeding in the Osprey Pandion haliaetus were studied during 1981–2006 in a small local population of about 20 pairs near the southern coast of Finland. The timing of breeding was estimated on the basis of the wing length of the largest young within the brood. The wing lengths were plotted against the date of the measurement to get a linear regression model characterizing the average developmental stage of broods in relation to date. This model was used when converting the measurements to correspond to the median date of all the measurements. Large-scale climatic phenomena were characterized by the winter and monthly indices of the North Atlantic oscillation (NAO). A significant positive relationship between the winter NAO index and timing of breeding in the Osprey, as indicated by the wing length of the largest young in the brood, was found. The relationship was even stronger when only the monthly value of January was considered. These relationships seemed to be linked with consequent earlier ice melting after mild winters. The results indicate prominent effects of large-scale climatic phenomena and that they can be monitored with relatively modest effort.     

Trophic interactions under climate fluctuations: the Atlantic puffin as an example

Co-occurrence in food requirements of offspring and food availability is a key factor determining breeding success. Prey availability is typically dependent on environmental conditions that are different from those influencing the predator''s decision regarding whether or not to initiate breeding, and is not always optimal at the peak of reproduction requirements. We investigated this relationship to understand better what determines the fledging success of the Atlantic puffin (Fratercula arctica). Colony data from Røst (northern Norway) covering a period of 27 years were analysed with parallel data on sea temperature and the size and abundance of the puffins'' main prey (the Norwegian spring-spawning herring, Clupea harengus). By fitting statistical models to the fledging success, we found that one effect of climate on this population of Atlantic puffins is indirect and mediated by sea temperature affecting the availability of first-year herring. The best model also demonstrates that the breeding success of the Røst puffins may be quantitatively predicted from the size of first-year herring and sea temperature.     

Ecological significance of 0-group fish in the Barents Sea ecosystem

Investigations into the 0-group fish in the Barents Sea have been carried out since 1965, with the goal of estimating the abundance of 0-group fish. 0-group abundance indices have been used in the assessment of the recruitment level and in recruitment variability studies. However, the ecological importance of the 0-group fish in the Barents Sea has been less studied. Although 0-group capelin, herring, cod and haddock are widely distributed in the Barents Sea, the central area seems to be the most important, accounting for approximately 50–80% of the annual biomass. The total biomass of the four most abundant 0-group fish species can be up to 3.3 million tonnes, with an average of 1.3 million tonnes (1993–2009). Wide distribution and high biomass of pelagically distributed 0-group fish make these fishes an important element in the energy transport between different trophic levels and different geographical areas, having a critical impact on the entire Barents Sea ecosystem. In recent years, capelin have shown a pronounced northward shift in biomass distribution, and several successive strong year classes occurred during warm temperature conditions. Cod biomasses were unexpectedly low during warm years and were positively correlated with spawning stock biomass, while the correlation with temperature was not significant. Haddock and herring show, as expected, increasing biomass with increased temperature when the spawning stock is at a sufficiently high level.     

Seasonal changes in age class structure and reproductive status of house mice on Marion Island (sub-Antarctic)

Feral house mice on sub-Antarctic Marion Island become reproductively active (males scrotal, females with perforate vaginas or pregnant) at an age >60 days and breed until death, which may occur at more than 13 months. Breeding is strongly seasonal; pregnant or lactating females were found only from October to May. A substantial proportion of mice old enough to breed in one summer overwinters to form a significant component of the breeding population the following summer but it is unlikely that any survive a second winter. The onset of breeding is closely synchronized with increasing day length but occurs about 2 months before mean temperature at the ground surface starts to increase significantly. Cessation of breeding is more closely associated with declining temperatures in late summer. For both males and females, the best correlation between reproductive activity and any of the temperature parameters measured was with average maximum temperature 1 cm above the ground. Competition for macroinvertebrate prey increases sharply in early winter due to high mouse numbers. The breeding season in 1991/1992 and 1992/1993 was at least 2 months longer than in 1979/1980, because the mice started breeding earlier, and stopped breeding later, in 1991/1993. The later cessation of breeding in 1991/1993 was despite the fact that there was a greater competition for macroinvertebrate prey, and that mean air temperatures during the early winter months were lower, than in 1979/1980.     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Meta-ecosystems and biological energy transport from ocean to coast: the ecological importance of herring migration

Ecosystems are not closed, but receive resource subsidies from other ecosystems. Energy, material and organisms are moved between systems by physical vectors, but migrating animals also transport resources between systems. We report on large scale energy transport from ocean to coast by a migrating fish population, the Norwegian spring-spawning (NSS) herring Clupea harengus. We observe a rapid body mass increase during parts of the annual, oceanic feeding migration and we use a bioenergetics model to quantify energy consumption. The model predicts strong seasonal variation in food consumption with a marked peak in late May to July. The copepod Calanus finmarchicus is the most important prey and 23×10⁶ tonnes (wet weight) of C. finmarchicus is consumed annually. The annual consumption-biomass ratio is 5.2. During the feeding migration 17% of consumed energy is converted to body mass. The biomass transported to the coast and left as reproductive output is estimated from gonad weight and is about 1.3×10⁶ tonnes for the current population. This transport is to our knowledge the world’s largest flux of energy caused by a single population. We demonstrate marked temporal variation in transport during the last century and discuss the effects of NSS herring in the ocean, as a major consumer, and at the coast, where eggs and larvae are important for coastal predators. In particular, we suggest that the rapid decline of lobster Homarus gammarus landings in Western Norway during the 1960s was related to the collapse of NSS herring. We also discuss spatial variation in energy transport caused by changed migration patterns. Both climate and fisheries probably triggered historical changes in the migration patterns of NSS herring. New migration routes emerge at the level of individuals, which in turn determines where resources are gathered and delivered, and therefore, how meta-ecosystems function.     

Predation as a probable mechanism relating winter weather to population dynamics in a North American porcupine population

An abundance index of an eastern Quebec population of North American porcupines (Erethizon dorsatum) has cycled with superimposed periodicities of 11 and 22 years from 1868 to 2000. This cycle closely followed 11- and 22-year cycles in solar irradiance and local weather (e.g., winter precipitation and spring temperature), generating the hypothesis that solar activity may affect porcupine abundance through effects on local weather. We investigated the mechanisms linking porcupine abundance to local weather conditions using a 6-year study (2000–2005) involving individual mark-recapture, radio tracking, seasonal survival analyses and identification of mortality causes. Summer (May–August) survival was high and constant over the study period, whereas winter (August–May) survival was lower and varied during the duration of our study. Variations in local winter precipitation explained 89% of the variation in winter survival. Porcupine predation rates appeared strongly related to snow conditions; 95% of depredated porcupines were killed when snow was covering the ground, and predation rates were higher in years with increased winter precipitation. Our data thus support the hypothesis that changes in predation rates under different snow conditions were the mechanism relating climate to porcupine population dynamics, via modifications of the local predator–prey interactions and impacts on porcupine winter survival. Our study adds to the growing body of evidence supporting an effect of climate on predator–prey processes. Also, it identifies one possible mechanism involved in the relationship between solar irradiance and porcupine population cycles observed at this study site over a 130-year period.     

Seasonal distribution and dive behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) of the White Sea–Barents Sea stock

Eight adult female harp seals (Pagophilus groenlandicus) of the White Sea–Barents Sea stock were tagged with satellite-linked dive recorders during the nursing period and followed from breeding in late February 1995 until moulting in late April 1995. Another ten adult harp seals of both sexes were tagged and followed from moult in early May 1996 until breeding in late February the following year. Between breeding and moult the seals were distributed along the coasts of Kola of Russia and eastern Finnmark of Norway, coinciding in time and space with the spawning capelin (Mallotus villosus). Between moulting and breeding they encircled the entire Barents Sea, mostly in open water, using the water column from 20 to 300 m, and in so doing by and large reflecting the annual migrations of the capelin. Capelin is therefore assumed to be the main source of prey for the White Sea–Barents Sea stock of harp seals, to be substituted, in part, by amphipods (e.g. Themisto libellula) in mid-summer and polar cod (Boreogadus saida) and herring (Clupea pallasii) in late autumn and winter. These data provide a baseline for the evaluation of the effects of future climatic change in the rich Barents Sea ecosystem.     

Relation between climatic factors,diet and reproductive parameters of Little Terns over a decade

We used 10 years of data on clutch size, egg size and diet, and 8 years of data on timing of laying on Little Terns (Sternula albifrons) breeding in Ria Formosa lagoon system, Algarve, Portugal to assess whether diet acts as an important intermediary between climatic conditions and breeding parameters. We used Generalized Linear Models to relate (1) the relative occurrence and size of the main prey species, sand smelts (Atherina spp.), with environmental variables, a large-scale climate variable, the North Atlantic Oscillation (NAO) index, and a local scale variable, the sea-surface temperature (SST), and (2) the respective effects of sand smelts relative occurrence, NAO index and SST on Little Tern breeding parameters. The diet of Little Terns was dominated by sand smelts, with a frequency occurrence of over 60% in all years. The winter SST (February) was negatively associated with the relative occurrence of sand smelts in the diet of Little Terns during the breeding season which, in turn, was positively associated with Little Tern clutch size. Our results suggest that negative NAO conditions in the Atlantic Ocean, often associated with rougher sea conditions (greater vertical mixing, stronger winds and lower SST) were related with earlier breeding, and lower SST in the surroundings of the colony during winter–spring favour the abundance of prey fish for Little Terns as well as their reproductive parameters. Climate patterns at both large and local scales are likely to change in the future, which may have important implications for estuarine seabirds in Southern Europe.     

Body mass and composition responses to short-term low energy intake are seasonally dependent in Steller sea lions (Eumetopias jubatus)

Steller sea lions (Eumetopias jubatus) were fed restricted iso-caloric amounts of Pacific herring (Clupea pallasi) or walleye pollock (Theragra chalcogramma) for 8–9 days, four times over the course of a year to investigate effects of season and prey composition on sea lion physiology. At these levels, the sea lions lost body mass at a significantly higher rate during winter (1.6 ± 0.14 kg day⁻¹), and at a lower rate during summer (1.2 ± 0.32 kg day⁻¹). Decreases in body fat mass and standard metabolic rates during the trials were similar throughout the seasons and for both diet types. The majority of the body mass that was lost when eating pollock derived from decreases in lipid mass, while a greater proportion of the mass lost when eating herring derived from decreases in lean tissue, except in the summer when the pattern was reversed. Metabolic depression was not observed during all trials despite the constant loss of body mass. Our study supports the hypothesis that restricted energy intake may be more critical to Steller sea lions in the winter months, and that the type of prey consumed (e.g., herring or pollock) may have seasonally specific effects on body mass and composition.     

Demographic responses of house mice to density and temperature on sub-Antarctic Marion Island

Recent changes in the climate of the sub-Antarctic may influence the number of house mouse (Mus musculus sensu lato) living on islands in the region. An increase in mouse numbers, as conditions became milder, could amplify the effects of climate change on native prey species. However, we have no direct evidence of the influence of climate on mouse numbers in the sub-Antarctic. We, therefore, assessed demographic trends in the mouse population on Marion Island between 1991 and 2001. Both the climate and mouse numbers were relatively stable during our study. Mice, however, increased their reproductive output in years when ambient temperatures were relatively high. Moreover, reduced reproductive output followed high densities at the onset of a breeding season, implying density-dependent limitation. We conclude that both temperature and density limited the increase in numbers during the summer breeding season. Major die-offs during winter probably limit population size and explain the relative stability in numbers across the 10 years of our study.     

Decline of a montane Mediterranean pied flycatcher Ficedula hypoleuca population in relation to climate

Certain populations of long‐distance migratory birds are suffering declines, which may be attributed to effects of climate change. In this article, we have analysed a long‐term (1991–2015) data set on a pied flycatcher Ficedula hypoleuca population breeding in nest‐boxes in a Mediterranean montane oak forest, exploring the trends in population size due to changes in nestling recruitment, female survival and female immigration. We have related these changes in population parameters to local climate, winter NAO index and to breeding density. During the last 25 yr the population has declined by half, mainly in association with a decrease in nestling mass and structural size which had repercussions on the probability of nestling recruitment to the population. Lower local nestling recruitment in certain years was linked to lower female immigration rate in the same years. On the other hand, the local survival of females remained stable throughout the study period. Laying date and breeding success were negatively affected by local temperatures while breeding, recruitment rate likewise by minimum temperature prior to breeding in April. As minimum April temperatures have increased across the study period, this may have affected recruitment and immigration rates negatively. On the other hand, tarsus length and body mass of nestlings were positively associated with winter NAO index, pointing to more global climatic links. Moreover, there was also a negative temporal trend in body mass of adults, implying increasingly difficult conditions for breeding. Declining recruit production in the study area could be attributed to a mismatch between the timing of arrival and breeding in the population, and the peak of food availability in this area.     

Density-dependence in the survival and reproduction of Bald Eagles: Linkages to Chum Salmon

During the late 20th Century, due to decreases in both contamination and persecution, bald eagle (Haliaeetus leucocephalus) populations increased dramatically. Currently, mechanisms regulating eagle populations are not well understood. To examine potential regulating processes in the Pacific Northwest, where eagles are no longer primarily regulated by contaminants or direct persecution, we examined bald eagle reproductive success, breeding populations, winter populations, mortality, and salmon stream use. Wintering and breeding eagle populations in south-coastal British Columbia (BC) quadrupled between the early 1980s and the late 1990s, and have since stabilized. Density-dependent declines in reproduction occurred during 1986–2009, but not through changes in site quality. Mid-winter survival was crucial as most mortality occurred then, and models showed that density-dependent reductions in population growth rates were partially due to reduced survival. Wintering eagles in British Columbia fed heavily on chum salmon (Oncorhynchus keta) runs, and then switched to birds in late winter, when mortality was highest. Eagles tended to arrive after the peak in salmon availability at streams in BC as part of a migration associated with salmon streams from Alaska to northern Washington. Eagles were most abundant in southern BC during cold Alaskan winters and in years of high chum salmon availability. We suggest that eagle populations in the Pacific Northwest are currently partially limited by density on the breeding grounds and partially by adult mortality in late winter, likely due to reduced late winter salmon stocks forcing eagles to exploit more marginal prey supplies. Larger eagle populations have affected some local prey populations. © 2011 The Wildlife Society.     

Effects of extrinsic and intrinsic factors on breeding success in a long lived seabird

There is growing concern over the impacts of climate change on animal species. Many studies have demonstrated impacts of climate change at the population level, and density dependent effects of climate are frequently reported. However, there is an increasing recognition of the differential impact of such factors on individuals since there is marked variation in individual performance. We investigated the relationships between breeding success and environmental conditions (winter NAO and one year lagged winter NAO) and intrinsic effects (colony size, pair bond duration, past breeding success rate) in the northern fulmar Fulmarus glacialis , using data from a long-term study commenced in 1950. There was a negative trend in breeding success over time, and a negative relationship with winter NAO and lagged winter NAO, which themselves had shown positive increases over the study period. The effects of lagged winter NAO remained after accounting for the linear trend. There was no evidence of density dependence, with breeding success positively related to colony size. We found strong evidence that breeding success was negatively related to pair bond duration but positively related to past breeding success rate. There was also an interaction between these two intrinsic effects such that those pairs that had historically been successful maintained success with increasing pair bond duration, whereas less successful pairs showed a decline. The prediction that there would be a differential impact of extrinsic factors among pairs was supported by an interaction between past breeding success rate and winter NAO, such that pairs with low past success rate exhibited a sharp decline in breeding success with increasing winter NAO, whereas more successful pairs did not. It is critically important to understand interactions between extrinsic factors and individual heterogeneity since a differential impact on individuals will affect population structure, and hence population dynamics.     

Diet of black guillemots and northern fulmars breeding beside a High Arctic polynya

In the Arctic, the availability of early-season open water (shore leads, polynyas) is a key factor influencing annual reproduction by marine birds, as these relatively productive locations provide the only sites where migrating and breeding birds can feed. We examined the diet of two marine birds, black guillemots (Cepphus grylle) and northern fulmars (Fulmarus glacialis), feeding in the Hell Gate–Cardigan Strait Polynya in the Canadian High Arctic during the breeding seasons of 1980–1984. There was little overlap in prey items consumed between these two species, except that both relied on similar sympagic amphipods. Guillemots fed principally on benthic prey, and compared to earlier High Arctic studies, they consumed proportionally more benthic fish, mysids and decapods. Fulmars appeared to feed on swarms of pelagic or surface prey, and in this polynya they consumed proportionally fewer fish and more polychaetes than has been reported for fulmars at other High Arctic locations. Given that climate change may alter marine food webs by changing the types and availability of prey, our results set a baseline of dietary information for these predators against which future conditions can be compared.